Trees, shrubs or herbs, monoecious, glabrous or with simple hairs; branching unspecialized
with spiral on distichous phyllotaxy, or phyllanthoid with leaves on main axes reduced to
cataphylls, subtending deciduous branchlets with distichous leaves. Leaves variable, simple,
symmetric or sometimes asymmetric, entire, penninerved, short-petioled; stipules varied, entire
to fimbriate, sometimes scarious along margins, often peltate, rarely hardening and becoming
spinescent on older branches. Inflorescences usually axillary, fasciculate or solitary, unisexual
or bisexual, sometimes flowers in racemes on panicles; bracteoles 2 per flower, entire to fimbriate
and often scarious along margins, often peltate. Male flowers: sepals 3 + 2 or 3 or sometimes 2
+ 2, free, imbricate, entire to fimbriate and often white on scarious along margins; petals absent;
disc glands of various shapes and sizes; stamens 3 - 6; filaments free or connate; anthers 2-
loculed, free or occasionally connate, vertically or transversely dehiscent; pistillode absent.
Female flowers: pedicellate or occasionally subsessile; sepals 3 + 2 or 3, as in male, occasionally
accrescent; petals absent; disc cupular-annular or flat, entire, crenate or lobed or divided into
segments; ovary usually 3-locular, in a few species 4 - 12-locular; locules biovulate; ovules
anatropous or amphitropous; styles 3, short, free or shortly connate, bifid. Fruits mostly
dehiscent capsules, occasionally baccate or drupaceous; cocci crustaceous, mostly 3, bivalved,
separating from a persistent columella; seeds 2 per locule, trigonous; testa smooth or sculptured,
exarillate; endosperms cartilaginous; embryo straight or slightly curved; cotyledons usually
broader than the radicle.
A large and very diverse genus of 750 - 850 species, widespread in tropical and subtropical
regions around the world; 51 species in India.
Literature.
CHAUDHARY, L. B. & R. R. RAO (2002). Taxonomic study of herbaceous species
of Phyllanthus L. (Euphorbiaceae) in India. Phytotaxonomy 2: 143 – 162. GANGOPADHYAY, M., T.
CHAKRABARTY & N. P. BALAKRISHNAN (2004). On the status of Phyllanthus airyshawii and P.
kozhikodensis (Euphorbiaceae). J. Econ. Taxon Bot. 28: 585 – 590. 2004 (2005). MITRA, R. L. & M.
SANJAPPA (2003). Phyllanthus parvifolius, P. clarkei (Euphorbiaceae) and related Indian taxa. Bull.
Bot. Surv. India 45: 1 – 20, ff. 1 – 4. 2003 (2004). WEBSTER, G. L. (1956 - 1958). A monographic study
of the West Indian species of Phyllanthus. J. Arnold Arbor. 37: 91 - 122, 217 - 268, 340 - 359; 38: 51 - 80,
170 - 198, 295 - 373; 39: 49 - 100, 111 - 212.
KEY TO THE SPECIES
| 1 a. Herbs, shrubs or trees; branching not phyllanthoid; leaves on main stems spiral or distichous,
not reduced to scales; stamens 3; filaments shortly connate for short length or free; anthers
dehiscing horizontally; styles bifid
|
3.
Isocladus
|
| b. Shrubs, herbs or trees; branching phyllanthoid, if not then plants shrubby; leaves on main
stems spirally arranged, leaves on deciduous branchlets distichous; stamens 2 - 15; filaments
connate for greater part; anthers dehiscing longitudinally or horizontally; styles entire or
bifid 2
|
2
|
| 2 a. Woody shrubs; sepals 4 in male, 6 in female, toothed, lacerate or fimbriate, rarely subentire;
stamens 2 or 3; filaments usually connate; pollen grains globose, porate; fruits capsular, dry
or somewhat fleshy
|
2.
Eriococcus
|
| b. Herbs, woody herbs or trees; sepals 5 or 6 in both male and female, entire, not lacerate nor
fimbriate; male sepals 5 or 6; stamens 2 - 15; filaments free or connate; pollen grains not
porate, areolate, colporate, striate or foveolate; fruits various
|
3
|
| 3 a. Stamens 2 - 15, free or more commonly united; pollen grains globose, areolate; carpels 3;
fruits 6-seeded
|
6.
Xylophylla
|
| b. Stamens 2 - 6; pollen grains colporate, striate or foveolate; carpels 3 or 4; fruits 6 or 8-seeded
|
4
|
| 4 a. Stamens 5 or 6; anthers elongate; pollen grains tricolporate; carpels 3 - 10; fruits baccate, 3-or 8 - 16-seeded
|
4.
Kirganelia
|
| b. Stamens 2 - 4; anthers short; pollen grains colporate, striate or foveolate; carpels 3 or 4; fruits
6-seeded
|
5
|
| 5 a. Trees with drupaceous or pithy fruits; pollen grains colporate; carpels 3 or 4
|
1.
Cicca
|
| b. Herbs or low woody shrubs with capsular fruits; pollen grains colporate, striate or
foveolate; carpels 3
|
5.
Phyllanthus
|
| 1 a. Fruits more or less fleshy, drupaceous or baccate
|
2
|
| b. Fruits dry dehiscent, capsular
|
6
|
| 2 a. Fruit a berry, 3.5 - 10 mm in diam.
|
3
|
| b. Fruit a drupe, 1 - 3 cm in diam.
|
4
|
| 3 a. Stamens of inner whorl connate at base; female pedicels filiform; styles 5 – 10 (-12), up to 0.5 mm long, triangular, free on very shortly connate at base; berries 5 – 10 (- 12)-locular
|
35.
Phyllanthus reticulatus
|
| b. Stamens all free; female pedicels thickened upwards; styles 3, 2 - 2.5 mm long, connate below
into a column (0.5 - 1 mm long), bifid above; berries 3-locular
|
33.
Phyllanthus glaucus
|
| 4 a. Leaves ovate, oblong to elliptic, 3 - 9 x 1.5 - 4 cm, distantly distichous; sepals 4; stamens 4; filaments free
|
1.
Phyllanthus acidus
|
| b. Leaves linear-oblong to narrowly oblong-elliptic, 3 - 22 x 1 - 7.5 mm, closely distichous;
sepals 6; stamens 3; filaments connate into a column
|
5
|
| 5 a. Leaves usually 100 or more pairs per branchlet, closely distichous and often over lapping, linear-oblong, 3 - 20 x 1 - 5 mm; petioles 0.4 - 1 mm long
|
2.
Phyllanthus emblica
|
| b. Leaves up to 50 pairs per branchlet, not closely distichous, not overlapping, only touching
each other, oblong, oblong-elliptic to linear-oblong, 10 - 25 x 2 - 10 mm; petioles 1 - 3 mm long
|
3.
Phyllanthus indofischeri
|
| b. Leaves always arising singly at leaf-axils
|
7
|
| b. Perennial undershrubs, shrubs or trees
|
20
|
| 8 a. Leaves white hispidulous beneath along margins; female flowers subsessile; fruits tuberculate on up to 0.5 mm long pedicels
|
51.
Phyllanthus urinaria
|
| b. Leaves beneath glabrous or at least not white hispidulous along margins; female flowers
pedicellate; fruits smooth, on more than 1 mm long pedicels
|
9
|
| 9 a. Proximal leaves of leaf-bearing branchlets double the size of the distal leaves
|
37.
Phyllanthus airyshawii
|
| b. All leaves of branchlets more or less of the same size
|
0
|
| b. Venation of leaves (at least lateral nerves) visible at least on the under surface; plants not
strandline
|
11
|
| 11 a. Cymules mostly (90%) consisting of one male and one female flowers
|
39.
Phyllanthus amarus
|
| 12 a. Stipules peltate, cordate or sagittate at base
|
13
|
| b. Stipules neither peltate nor cordate or sagittate at base
|
15
|
| b. Anthers free, transversely dehiscent; ovary and fruits verrucose or muriculate
|
14
|
| 14 a. Leaves 5 - 10 mm long, thickened along margins; male flowers subsessile; female pedicels 1.5
- 3 mm long; fruiting pedicels 3 - 4 mm long
|
28.
Phyllanthus narayanswamii
|
| b. Leaves 6 - 25 mm long, flat along margins; male pedicels 1.5 - 3 mm long; female pedicels (2.5
-) 4 - 10 mm long; fruiting pedicels 5 - 15 mm long
|
32.
Phyllanthus simplex
|
| 15 a. Sepals 5; stamens 5
|
16
|
| b. Sepals 6; stamens 3
|
17
|
| 16 a. Herbs, becoming shrubby, up to 45 cm high; cataphylls present; stipules linear to triangular
lanceolate, entire, unlobed; leaves elliptic to oblong-elliptic, lateral veins distinct and raised
on both surfaces; female disc annular, entire, fleshy
|
36.
Phyllanthus tenellus
|
| b. Herbs, up to 10 cm high; cataphylls absent; stipules linear-lanceolate, often lobed or laciniate
at basal portion; leaves spathulate or obovate, lateral veins not visible; female disc of 5
distinct rectangular or tongue-shaped glands
|
38.
Phyllanthus ajmerianus
|
| 17 a. Leaves scabridulous with white hairs on lower surface; venation distinct on the upper surface;
stipules serrate
|
31.
Phyllanthus scabrifolius
|
| b. Leaves not scabridulous nor white hairy beneath; venation inconspicuous on the upper
surface; stipules entire
|
18
|
| 18 a. Female disc crenate or shallowly lobed; leaves usually narrowed at apex to a point; anther-
thecae dehiscing horizontally or obliquely
|
42.
Phyllanthus debilis
|
| b. Female disc deeply dissected; leaves rounded, obtuse, apiculate or subacute at apex; antherthecae
dehiscing vertically
|
19
|
| 19 a. Branchlets usually scabrid-hispidulous at the angles; petioles 0.4 - 1 mm long; stipules 0.4 -
1 mm long; bracts up to 1 mm long; filaments united into a column
|
43.
Phyllanthus fraternus
|
| b. Branchlets glabrous; petioles 1 - 1.5 mm long; stipules 1 - 2 mm long; bracts 1 - 2 mm long;
filaments free at least halfway above
|
30.
Phyllanthus rheedei
|
| 20 a. Sepals in male flowers 4 (Eriococcus)
|
21
|
| b. Sepals in male flowers 5 - 6
|
37
|
| b. Fruits not enclosed by the enlarged calyx
|
22
|
| 22 a. Branchlets leafless or with suppressed leaves along the upper portion
|
23
|
| b. Branchlets with normal leaves at upper portion
|
26
|
| 23 a. Branchlets and often nerves on leaves beneath pubescent
|
24
|
| b. Plants entirely glabrous
|
25
|
| 24 a. Leaves 1 - 3 x 0.5 - 1.3 cm, chartaceous to thinly coriaceous, glabrous; lateral nerves 5 or 6
pairs; petioles ca 1 mm long; male cymules 1-flowered; filaments almost free; fruits glabrous
|
14.
Phyllanthus pulcher
|
| b. Leaves 2 - 5 x 1 - 3 cm, membranous to thinly chartaceous, mostly pilose beneath on nerves;
lateral nerves 6 - 11 pairs; petioles 1.5 - 3 mm long; male cymules 3 - 5- flowered; filaments
united into a column; fruits evanescently papillose-puberulous
|
16.
Phyllanthus sikkimensis
|
| 25 a. Apices of branchlets leafless; leaves acute to attenuate-mucronate at apex, often marbled
with white, purple or brown along veins; stipules epeltate; male sepals entire; disc 2-lobed,
fleshy; fruits 6 - 8 mm in diam.
|
4.
Phyllanthus baillonianus
|
| b. Apices of branchlets with suppressed leaves; leaves apiculate at apex, not marbled; stipules
peltate; male sepals lanceolate; disc of 4 distinct glands; fruits ca 10 mm in diam.
|
9.
Phyllanthus gageanus
|
| 26 a. Leaves 3 - 12 cm long
|
27
|
| b. Leaves 0.5 - 3 cm long
|
31
|
| 27 a. Branchlets tomentellous
|
28
|
| b. Plants entirely glabrous
|
29
|
| 28 a. Leaves acute, subacute or shortly acuminate (acumen up to 5 mm long) at apex; female
cymules pedunculate, fasciculate; female and fruiting pedicels 10 - 15 mm long; fruits 4 - 6
mm in diam.
|
19.
Phyllanthus tetrandrus
|
| b. Leaves usually acuminate or caudate (acumen on cauda 10 - 20 mm long) at apex; female
cymules epedunculate, 1-flowered; female and fruiting pedicels 2 - 7 mm long; fruits 8 - 20
mm in diam.
|
10.
Phyllanthus gomphocarpus
|
| b. Sepals (at least in male flowers) fimbriate or denticulate; male pedicels up to 5 mm long
|
30
|
| 31 a. Leaf bearing branchlets arising in fascicles of 3 - 5; leaves linear-oblong to narrowly oblong
|
18.
Phyllanthus talbotii
|
| b. Leaf bearing branchlets arising singly; leaves otherwise
|
32
|
| 33 a. Leaves 0.5 - 1 cm long; male cymules 1-flowered; anthers subsessile; female and fruiting
pedicels up to 10 mm long
|
6.
Phyllanthus chandrabosei
|
| b. Leaves 1 - 2.7 cm long; male cymules 2 or 3-flowered; anthers on ca 1 mm long column;
female and fruiting pedicels 10 - 20 mm long
|
7.
Phyllanthus cinereus
|
| 34 a. Lateral nerves of leaves immersed or faint; nervules hidden; stipules peltate, hastate at base
|
35
|
| b. Lateral nerves and nervules of leaves visible at least on the under surface; stipules epeltate,
not hastate at base
|
36
|
| 35 a. Phyllanthoid branches crowded towards tip of stem; lateral nerves of leaves hidden; sepals
lacerate; bracts lacerate to denticulate
|
17.
Phyllanthus singampattianus
|
| b. Phyllanthoid branches scattered along the stem; lateral nerves of leaves faint; sepals entire to
undulate; bracts entire
|
15.
Phyllanthus rangachariarii
|
| 36 a. Branchlets rufous tomentellous to scattered whitish puberulous or pilose; lateral nerves of leaves 5 or 6 pairs, arising at 35° to 45° angles with the midrib; filaments ca 0.3 mm long, almost free; fruits without a stipe
|
14.
Phyllanthus pulcher
|
| b. Branchlets glabrous; lateral nerves of leaves 6 - 11 pairs, arising at 45° to 70° angles with the midrib; filaments ca 1 mm long, connate into a column; fruits shortly stipitate
|
11.
Phyllanthus heyneanus
|
| 37 a. Branchlets leafless and bearing dense-flowered fascicles at apical portion; staminal column
long, slender, exserted; stylar column stout, exserted
|
38
|
| b. Branchlets with normal leaves at apical portion
|
39
|
| 38 a. Shrubs or small trees, 2 – 3 m tall; leaf-bearing branchlets 5 – 17 cm long, glabrous; petioles
glabrous; pseudoracemes 1 – 5 cm long; fruiting pedicels 3 – 4 mm long
|
21.
Phyllanthus andamanicus
|
| b. Trees 3 – 12 m tall; leaf-bearing branchlets 15 – 60 cm long, rufous tomentellous; petioles
rusty tomentellous; pseudoracemes 10 – 20 cm long; fruiting pedicels 5 – 10 mm long
|
23.
Phyllanthus columnaris
|
| b. Inflorescences axillary, fasciculate or solitary or flowers in axillary or supra-axillary racemes
or spikes
|
40
|
| b. Flowers in axillary fascicles or solitary
|
41
|
| 41 a. Stipules peltate
|
42
|
| b. Leaves glabrous on the lower surface
|
43
|
| 43 a. Male pedicels ca 15 mm long; sepals recurved; filaments 2 - 3.5 mm long, united below (for
3/4th length) into a slender column
|
44.
Phyllanthus griffithii
|
| b. Male pedicels 1.5 - 3 mm long; sepals erect or spreading; filaments up to 0.8 mm long, free or
shortly connate below into a column
|
44
|
| b. Female and fruiting pedicels 5 - 12 mm long
|
45
|
| 45 a. Leaves usually obovate, rounded or subobtuse at apex, membranous to chartaceous; flowers
arising on short pulvini; fruits 3 - 3.5 mm in diam.
|
46.
Phyllanthus praetervisus
|
| b. Leaves narrowly oblong, elliptic to ovate-elliptic, apiculate or acute-mucronate at apex,
coriaceous to chartaceous; flowers arising amidst tufts of bracts, often forming short
brachyblasts; fruits 2 - 2.5 mm in diam.
|
48.
Phyllanthus roeperianus
|
| b. Leaves 0.2 - 3.5 cm long
|
47
|
| 47 a. Stipules on main branches hardening and becoming spiniform
|
48
|
| b. Stipules on main branches not modified into spines, or if spiniform then deciduous
|
49
|
| 48 a. Stipular spines persistent; leaves oblong, elliptic-oblong to linear-oblong, 2 – 8 x 1 - 3 mm,
stiff chartaceous; filaments connate into a column to about the middle; styles ca 1.5 mm long,
united below into a column; fruiting pedicels up to 1.5 mm long
|
24.
Phyllanthus lawii
|
| b. Stipular spines deciduous; leaves obovate (or narrowly so) obovate-elliptic to obovate-
oblong 4 - 15 x 2 - 7 mm, membranous; filaments free; styles 0.5 - 1 mm long, free; fruiting
pedicels 2 - 6 mm long
|
47.
Phyllanthus pseudoparvifolius
|
| 49 a. Leaves linear-oblong to narrowly oblong, usually slightly constricted at the middle; nervules
very prominent on the under surface
|
29.
Phyllanthus polyphyllus
|
| b. Leaves obovate, elliptic to oblong, not constricted at the middle; nervules obscure or less
conspicuous on the under surface
|
50
|
| 50 a. Petioles 0.2 - 1 mm long
|
51
|
| b. Petioles 1 - 2.5 mm long
|
52
|
| 51 a. Branchlets scaberulous at angles; leaves membranous to firmly chartaceous, rounded, truncate,
retuse or apiculate at apex; lateral nerves prominent to very prominent on the under surface;
petioles 0.5 - 1 mm long; styles free
|
41.
Phyllanthus clarkei
|
| b. Branchlets glabrous; leaves coriaceous, obtuse-mucronate at apex; lateral nerves faint on the
under surface; petioles 0.2 - 0.4 mm long; styles connate below into a column
|
50.
Phyllanthus sanjappae
|
| 52 a. Plants densely hairy; leaf-bearing branches solitary at each node; leaves firmly chartaceous,
curling down along margins; flowers subtended by tufts of subflabellate fimbriate bracts;
male flowers solitary or accompanied by a female flower in cymules
|
40.
Phyllanthus brevipes
|
| b. Plants entirely glabrous; leaf-bearing branches often 2 or 3 at each node; leaves membranous,
flat along margins; flowers subtended by entire bracts; male flowers paired or up to 10-
flowered in each cymule
|
45.
Phyllanthus leschenaultii
|
